The fauna of Guatemala and Belize: survey and biogeographical affinities

Alma Estrella García-Morales & Manuel Elías-Gutiérrez El Colegio de la Frontera Sur. Av. Centenario km 5.5 Chetumal 77900, Quintana Roo, México; [email protected]

Received 11-Xii-2001. Corrected 30-viii-2006. Accepted 15-Xii-2006.

Abstract: Rotifer samples were obtained from 29 localities in northern Guatemala and central-southern Belize during March 2000 and June 2001. A total of 119 species were identified. Ten selected taxa are illustrated and commented: Euchlanis semicarinata, Lepadella apsicora, L. cryphaea, Lecane curvicornis f. lofuana, L. whitfordi, Monommata maculata, Scaridium bostjani, Trichocerca elongata f. braziliensis, and T. hollaerti. The species Lepadella rhomboidula is a first record for the American Continent. The species are 71 % cosmopolitan, 6 % tropicopolitan, and 4.2 % restricted to the subtropics. The Guatemala species number range was Petén-Itza lake (53 taxa), and Raxruja pool (three). La Democracia pool (49 taxa), and the Blue Hole sink-hole (six spe- cies) were the extremes in Belize. In total, 68 of the recorded taxa are new for Guatemala and 91 for Belize. Additionally, 47 species are registered by the first time in Central America. A comparison between these two countries and Mexico revealed that the south part of the latter conform a cluster with them, emphasizing the transitional character of this region between the Nearctics and the Neotropics. Furthermore, Guatemala and Belize have differences in species assemblages, as a response to the nature of their particular environments and topographical accidents. Rev. Biol. Trop. 55 (2): 569-584. Epub 2007 June, 29.

Key words: rotifera, , diversity, distribution, Guatemala, Belize.

In Central America, studies on taxonomy Elías-Gutiérrez (2004) recorded 128 species and distribution of are relatively scarce from 36 localities in the south of Mexico. A and old (ie. Collado et al. 1984). Currently, previous study by Sarma and Elías-Gutiérrez knowledge on the rotifer fauna of Guatemala is (1999a) in 12 localities from Yucatan rendered poor compared with North and South America 102 species. In both papers, a total of 37 new (Koste and De Paggi 1982, De Paggi and records for the region were figured, majority Koste 1995, Segers 1997, Sarma 1999, Sarma of them regarded as warm-stenotherm (García- and Elías-Gutiérrez 1999a, b, García-Morales Morales and Elías-Gutiérrez 2004). and Elías-Gutiérrez 2004). On the other hand, North of Guatemala and Belize possess Belize’s fauna is completely unknown; there highly contrasting environments due to topo- are no previous reports on the rotifers from graphic accidents, quite different with the this country. Lately, a preliminary study on homogeneous adjacent south of Yucatan Cladoceran fauna published by Elías-Gutiérrez Peninsula. Nevertheless, all of them belong to et al. (2006), emphasized about the diverse the same biotic province, named Campechano- aquatic systems found in this region, and the Petenense (UNAM 1990). As a result of the effect on their fauna. In case of rotifers, recent presence of mountain systems (for example publications are limited to the Mexican side, Mayan mountains in Belize and Sierra Chinajá close to both countries. García-Morales and in north of Guatemala) and other topographic

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 55 (2): 569-584, June 2007 569 accidents, both countries have diverse micro- geographical coordinates, and environmental habitats reflected in the vegetation and aquatic data of each sampling site are resumed in Table systems. As a result, in the case of fish fauna for 1. were sorted with a stereomicroscope example, 13 endemic species dwell in north of and identified using a compound microscope. Guatemala (Valdéz-Moreno et al. 2005). Many Identification of rotifers was made following of the aquatic systems from these regions still Koste 1978, Nogrady et al. 1995, Segers 1995, remain as pristine and non polluted environ- De Smet and Pourriot 1997, Nogrady and ments (Valdéz-Moreno et al. 2005, pers. obs.). Segers 2004. Also we consulted the Academy The aims of this study were to provide of Natural Sciences of Philadelphia database on data on the species of the rotifer fauna found rotifers (Jersabek et al. 2003). When needed, in zooplankton collections gathered in several animals were dissolved in a sodium hypoclo- freshwater bodies from Guatemala and Belize, rite solution to allow examination of trophi. to include a comment about the distribution of Information about varieties and forms is pro- these animals in both countries, and to compare vided because rotifer taxonomy is still under the species assemblage with adjacent Mexico. discussion. For example, former varieties have been recently proposed to be good species (ie. Sarma and Nandini 2002 proposed to consider MATERIALS AND METHODS macracanthus, a former variety of Brachionus patulus, as a different species). Selected new Rotifers were qualitatively collected using records in Central America were illustrated standard plankton net; 50 µm mesh size, attached by using a camera lucida. All material and to a handle in the littoral zone. Horizontal and original samples are deposited at the reference vertical tows, with the same kind of net were performed, when a limnetic zone (more than 5 m deep) was clearly defined in the surveyed water body. As a minimum, 200 l of water were filtered for each sample. After collections, mate- rial was preserved in 4 % formalin. At least five samples represented bigger systems, as Petén Lake, and the small puddles were characterized by one. Littoral sampling included all possible habitats for rotifers: submersed vegetation, bot- tom dwellers, etc. Temperature, depth, and Secchi transparency were measured at every site to give a reference about the environment where the rotifers dwell. In total, 11 localities were sampled in Guatemala and 18 from Belize (Fig. 1). Each place was visited once. Data on physical parameters and geographical coordi- nates of the studied sites are presented in Table Fig. 1. Guatemala and Belize with localities collected 1. In general, the surveyed systems are shallow (some of them were grouped due to their proximity). 1. (0.2-1.5 m depth), except for Petén-Itza Lake, Charco Seco; 2. Laguna and Estanque Crooked Tree; 3. the largest system. Transparency showed values Sand Hill; 4. La Democracia and Dos Charcos; 5. Charco ranging between 0.17-2 m, whereas tempera- Dos Puentes and Spanish Lockout I-III; 6. Blue Hole and ture had values between 23-37 °C, ranges also Caves Branch; 7. Pine Ridge I-III; 8. Río on Pools and Río Puente Largo; 9. Caracol; 10. Petén-Itza Lake; 11. Charco found in other surveys from the region (Sarma El Pato and Sayaxché; 12. Charco Raxruja, Río Candelaria, and Elías-Gutiérrez 1999a, García-Morales Charco Rayulia and Río San Simón; 13. Charco Sebol and and Elías-Gutiérrez 2004). Date of collection, Río Sebol; 14. Río Chajmaic and Río Dolores.

570 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 55 (2): 569-584, June 2007 TABLE 1 Physical variables and location of the water bodies sampled

Number Collecting Temperature Maximum Secchi Place Geographical coordinates of samples date (°C) depth (m) disk (m) GUATEMALA Latitude N Longitude W A Laguna Petén-Itza 16°55’44” 89°53’38” 5 21/03/00 30 40 2 B Sayaxché 16°12’33.6” 90°10’47.3” 1 18/05/01 32 1 0.25 C Charco Raxruja 15°53’29” 90°05’51” 1 27/03/00 29 0.39 0.39 D Río Candelaria 15°53’04” 90°11’12” 2 27/03/00 28 0.5 0.5 E Charco Sebol 15°40’25” 89°56’39” 3 23/03/00 26 0.8 0.8 F Río Sebol 15°40’25” 89°56’39” 2 22/03/00 ND 1 1 G Charco Río Chajmaic 15°43’15” 89°56’24” 1 23/03/00 23 0.4 0.4 H Río Dolores 15°44’17” 90°22’34” 3 26/03/00 26 0.5 0.5 I Charco Rayulia 15°54’41” 90°06’06” 1 27/03/00 29 1.17 1.17 J Río San Simón 15°50’39” 90°17’01” 1 25/03/00 25 0.25 0.25 K Charco del Pato 16°07’36” 90°10’53” 1 27/03/00 29 0.79 0.47 BELIZE L Charco Seco 17°55’31.5” 88°31’10” 1 14/06/01 36 0.2 0.2 M Charco Dos Puentes 17°13’27.6” 88°52’29.2” 1 14/06/01 34 1.5 1.5 N Spanish Lokcout I 17°13’54.1” 88°55’49.2” 1 14/06/01 36 0.2 0.2 Ñ Spanish Lokcout II 17°13’59.4” 88°56’04.4” 1 14/06/01 35 0.55 0.41 O Spanish Lokcout III 17°14’17.1” 88°56’49.3” 1 14/06/01 37 0.2 0.2 P Pine Ridge I 17°02’55.6” 88°56’57.8” 1 15/06/01 25.5 0.56 0.56 Q Pine Ridge II 17°03’04.9” 88°57’17.5” 1 15/06/01 25 0.33 0.33 R Pine Ridge III 17°02’56.1” 88°57’03.1” 1 15/06/01 26 0.4 0.4 S Caracol 16°45’34.1” 89°07’13.7” 1 15/06/01 28 0.5 0.28 T Laguna Crooked Tree 17°46’30.4” 88°31’51.5” 2 16/06/01 30 0.7 0.7 U Estanque Crooked Tree 17°46’24.7” 88°30’54.8” 1 16/06/01 32 0.73 0.73 V Blue Hole 17°09’26.6” 88°41’00.6” 2 16/06/01 25 0.56 0.56 W Rio on Pools 16°59’16.6” 88°58’40.3” 1 15/06/01 30 0.5 0.5 X Río Puente Largo 16°51’52.6” 89°02’21” 2 15/06/01 32 0.48 0.175 Y La Democracia 17°21’53.9” 88°32’20.7” 1 16/06/01 29 0.22 0.22 Z Sand Hill 17°35’41.5” 88°21’40” 1 16/06/01 35 0.25 0.25 AA Dos Charcos 17°21’53.9” 88°32’20.7” 1 16/06/01 30 0.36 0.36 AB Caves Branch 17°08’53.2” 88°42’33” 1 16/06/01 28 0.67 0.67 collection located in El Colegio de la Frontera ence/absence) (Legendre and Legendre 2000) Sur, Chetumal Unit. between all localities with known rotifer fauna. To compare the fauna between Belize and In the case of Mexico, we used already published Guatemala as well as Mexico, we performed a data by Silva-Briano and Segers (1992), Rico- cluster analyses with the Sorensen Index (pres- Martínez and Silva-Briano (1993), Kutikova

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 55 (2): 569-584, June 2007 571 and Silva-Briano (1994), Sarma et al. (1996, and possibly Europe (Koste 1978, Segers and 2000), Sarma and Elías-Gutiérrez (1997, 1998, Sarma 1993, Jersabek et al. 2003). 1999a, b, 2000), Silva-Briano and Adabache- Lepadella rhomboidula (Bryce, 1890) Ortíz (1999), Rico-Martínez et al. (2003) and (Fig. 5): Known by some as L. triptera f. García-Morales and Elías-Gutiérrez (2004). rhomboidula (see Koste 1978). In general view, Due to the fact that Mexico has a bigger size body rhomboid. With a median keel on dorsal with a complex topography, the records of plate. Ventral margin of head “U”-like. Europe, rotifer fauna were grouped as Central Plateau, India, Indonesia, and New Zealand. This is the Pacific slope, North, Southeast and Yucatan first record of this species in the Americas, it Peninsula regions. was found only in one locality from Belize Mayan Mountains. Lecane curvicornis f. lofuana (Murray, RESULTS 1913) (Fig. 6a, b): Ventral plate broader than dorsal, two antero-lateral spines. Dorsal and A total of 119 species were recorded ventral margins slightly convex. Posterior end (Table 2); 68 are new records to Guatemala, of lorica trilobate. The species is relatively com- and 91 taxa are recorded by the first time in mon, with cosmopolitan distribution (Segers Belize. The species found belong to two orders, 1995), but the f. lofuana has been reported more 20 families, 35 genera and eight forms. The rarely, basically from Singapore and Brazil. highest number of species was contained in Lecane whitfordi (Ahlstrom, 1938) (Fig. Lecane, with 37 species (31 %), Trichocerca 14 7a, b): Related with L. obtusa. Dorsal plate (11.8 %), and Lepadella with 11 taxa (9.32 %). broader than ventral, head aperture margins These results agree in part with other reports straight or slightly convex. Antero-lateral cor- from tropical zones, where Lecane is the most ners with long triangular spines. Foot simple common genus, followed by Lepadella (Segers with a toe ending in two separated claws. et al. 1998, Sarma and Elías-Gutiérrez 1999a). Known only from two localities in North In the following section, a short descrip- Carolina (USA), Lake Waccamaw, the type tion of selected taxa considered as new records locality (Segers 1995), and Lake Mattamuskeet for Central America is provided. Distributional (Jersabek et al. 2003). Possibly a record of L. data follow De Ridder and Segers 1992, unless obtusa by Paggi (1989) could be L. whitfordi, otherwise stated. according to Segers (1995). It was found only Euchlanis semicarinata Segers, 1993 (Fig. in one locality from Belize. 2): With a characteristic shallow dorsal keel. Monommata maculata Harring and Dorsal and ventral plates present. Nearly cir- Myers, 1924 (Fig. 8a, b): Body slender. Foot cular in transversal view. Known from Nigeria, with two segments, toes unequal in length. French Guyana, and Bolivia (Pourriot 1996, Mastax with fulcrum conical, short; rami with Segers et al. 1998). complex dentition. Unci with three long teeth. Lepadella apsicora Myers, 1934 (Fig. 3a, Manubrium curved, with lamella. A com- b): Body ovoid, with a characteristic “V”- mon species with cosmopolitan distribution shaped sinus. Distal segment of foot longer (Nogrady et al. 1995). than proximal. Toes asymmetric. Known from Scaridium botsjani Dames and Dumont, USA and Ecuador (Myers 1934, Koste and 1974 (Fig. 9): Trunk broad. Second segment Shiel 1989, Jersabek et al. 2003). of foot long. Toes relatively long. Fulcrum Lepadella cryphaea Harring, 1916 (Fig. with a median-ventral crest. Basal plate well 4a, b): Related with L. acuminata. Lorica ovoid, developed. Rami with long teeth. Manubria posterior dorsal plate pointed. Characterized by relatively narrow, with rounded alulae. Nepal the presence of folds and one dorsal-medi- is the type locality, but it is considered a warm an keel. Known from Canada, USA, Brazil stenoterm in all tropics (Segers 1995).

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------L ------K ------J X X X I ------X X TABLE 2 TABLE ------H X X X ------G X X X X X X X List of species and localities ------F X ------E X X X X X ------D X X X ------C X X ------B X X X X ------A X X X X X X X X X X X •

• • (Daday, 1905) * (Daday, (Gosse, 1886) + (Gosse, 1886) * + Rousselet, 1911 Rousselet, * 1911 De Beauchamp, 1907 (Levander, 1894) (Levander, * + (Ehrenberg, (Ehrenberg, 1832) * + Carlin, 1943 * Ehrenberg, Ehrenberg, 1832 * + (Lucks, 1912) (Hermann, 1783) * (Gosse, 1851) + Segers, 1993 (Gosse, 1851) (Ehrenberg, (Ehrenberg, 1832) * + (O. F. Müller,1786) * Müller,1786) (O. F. f. macracanthus (Apstein, 1907) + Gosse, 1887 +

Carlin, 1939 * + (Hauer, 1953) (Hauer, * Proalides tentaculatus Proalides Anuraeopsis fissa Brachionus havanensis B. patulus B. patulus B. quadridentatus Keratella americana K. cochlearis K. lenzi K. tropica K. tropica Platyias quadricornis Manfredium eudactylota Manfredium Dipleuchlanis propatula Dipleuchlanis propatula Euchlanis dilatata E. incisa E. oropha E. oropha E. semicarinata Tripleuchlanis plicata Tripleuchlanis Mytilina bisulcata M. ventralis Species Order Order Ploimida 1. Family Epiphanidae 2. Family Brachionidae 3. Family 4. Family Mytilinidae 5. Family Trichotriidae

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 55 (2): 569-584, June 2007 573 ------X X X X X X AB ------X X X X X AA ------Z X X X X X X ------Y X X X X X X X X X X X ------X X X X X X X ------X X X W ------V X ------U X X X X ------T X X X ------S X X X X X ------R X X ------Q X X X ------P X X ------O X X ------Ñ X X X X ------N X ------X X X X M ------L X X ------K X X X ------J X X X X I ------X ------H X X X X X ------G X X X X X X TABLE 2 (Continued) TABLE List of species and localities ------F X X ------E X X X X X X ------D X X X X ------C X X ------B ------A X X X X X X X X X X X • • •

+ • • • •

• (Ehrenberg, (Ehrenberg, 1832) * + • (Murray, 1913) * (Murray, (Gosse, 1867) * + (Ehrenberg, (Ehrenberg, 1834) • (Ehrenberg, (Ehrenberg, 1831) (Ehrenberg, (Ehrenberg, 1830) * (Stenroos, 1898) (Schmarda, 1859) * + (Bryce, 1890) (Gosse, 1886) * + Harring, 1914 * + bicuspidata (Perty, 1850) (Perty,

Harring, 1916 Myers, 1934 (O. F. Müller, 1786) Müller, * (O. + F. (Thorpe, 1893) (Ehrenberg, (Ehrenberg, 1830) * Harring, 1914 * + (Stenroos, 1898) (Gosse, 1886) * + (O. F. Müller, 1786) Müller, * (O. F. (O. F. Müller, 1786) Müller, * (O. + F. (Gosse, 1851) * + Myers, 1934 Trichotria tetractis Trichotria Macrochaetus collinsi Macrochaetus M. sericus Colurella adriatica Colurella C. obtusa C. sulcata C. uncinata f. Lepadella acuminata L. apsicora L. cryphaea L. ehrenbergi L. ehrenbergi L. ovalis L. patella L. quadricarinata L. rhomboides L. triba L. triptera L. rhomboidula Lecane arcula Lecane arcula L. bulla L. closterocerca L. closterocerca L. cornuta L. crepida L. crepida L. curvicornis f. nitida Species 6. Family Colurellidae 7. Family Lecanidae

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• • •

• • (Murray, 1913) (Murray, Harring, 1914 * (Ehrenberg, (Ehrenberg, 1832) * + Segers, 1991 + (Ehrenberg, (Ehrenberg, 1834) * + Edmondson, 1935 * (Daday, 1905) * (Daday, + (Daday, 1897) * (Daday, Harring & Myers, 1926 (Murray, 1913) + (Murray, (Jennings, 1896) * + (Murray, 1913) + (Murray, Hauer, 1929 Hauer, * (Eckstein, 1883) * + f. ercodes f. ercodes (Turner, 1892) * (Turner, + Harring, 1914 *

(Stokes, 1896) * + (Bryce, 1892) * (Murray, 1913) * (Murray, + Harring & Myers, 1926 * (Murray, 1913) * (Murray, (Ehrenberg, (Ehrenberg, 1832) * Harring & Myers, 1926 (Harring, 1914) * (Murray, 1913) * (Murray, + (Murray, 1913) * (Murray, + (O. F. Müller, 1776) Müller, * (O. + F. L. curvicornis f. lofuana L. decipiens L. doryssa L. furcata L. furcata L. haliclysta L. hamata L. hastata L. hornemanni L. inermis L. leontina L. ludwigii L. ludwigii L. luna L. lunaris L. margarethae L. margarethae L. monostyla L. nana L. obtusa L. papuana L. pertica L. pyriformis L. quadridentata L. rhenana L. rhytida L. rugosa L. signifera L. spinulifera Species

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(Murray, 1913)* + (Murray, • Ehrenberg, Ehrenberg, 1840* (Ehrenberg, (Ehrenberg, 1832)* + Harring & Myers, 1924* (Gosse, 1887)* (Gosse, 1886)* Harring & Myers, 1922+ Dames & Dumont, 1974* + Dujardin, 1841* + Ehrenberg, Ehrenberg, 1832* Myers, 1930* (Ahlstrom, 1938) * (Meissner, 1908) (Meissner, * + (Wierzejski & (Wierzejski Zacharias, 1893)+ (Gosse, 1887) * + De Smet, 1990* (Rousselet, 1896)* Harring & Myers, 1926 * Ehrenberg, Ehrenberg, 1834* (Lucks, 1912)* (Ehrenberg, (Ehrenberg, 1838)* + Yamamoto, 1951 * Yamamoto, L. stenroosi L. stenroosi L. subtilis L. uenoi L. ungulata L. whitfordi L. whitfordi Lindia torulosa Eosphora anthadis Cephalodella forficula C. gibba Monommata maculata M. cf. phoxa Notommata cerberus N. copeus N. saccigera Scaridium botsjani Tetrasiphon hydrocora hydrocora Tetrasiphon Trichocerca bicristata Trichocerca T. bidens T. T. capucina T. T. collaris T. T. elongata f. T. braziliensis T. flagellata Hauer, 1937* flagellata T. Hauer, T. hollaerti T. Species 8. Family Lindiidae 9. Family Notommatidae 10. Family Trichocercidae

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• • •

• • • • + I delson, 1925+ + •

Harring, 1913* • • (Ehrenberg, (Ehrenberg, 1832)* Hauer, 1926* Hauer, + Gosse, 1887+ (Bergendal, (Bergendal, 1892)+ (De Guerne, 1888)+ Weisse, 1848 Weisse, Carlin, 1943* + Harring & Myers, 1928+ Hauer, 1938* Hauer, (Leydig, 1854)* + (Gosse, 1886)* + (Laterbourn, 1898)* + (Jennings, 1903) (Gosse, 1851)* (Hauer, 1931)* (Hauer, (Wierzejski, 1893)* (Wierzejski, + (Hermann,1783)* + (Gosse, 1887)+ T. iernis T. T. myersi T. T. pusilla T. T. similis T. T. stylata T. T. tenuior T. T. weberi T. ovalis Polyarthra cf. dolichoptera P. cf. vulgaris P. Synchaeta longipes Asplanchna girodi Asplanchna girodi A. sieboldi Asplanchnopus hyalinus Dicranophorus grandis D. prionacis Testudinella ahlstromi ahlstromi Testudinella T. amphora T. T. patina T. Limnias melicerta Species 11. Family 11. Gastropodidae 12. Family Synchaetidae 13. Family Asplanchnidae 13. Family 14. Familia Dicranophoridae Order Order Gnesiotrocha 15. Family Testudinellidae 16. Family

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 55 (2): 569-584, June 2007 577 - - - - - Trichocerca elongata braziliensis (Murray, AB 1913) (Fig. 10): Body cylindrical. Anterior - - - - - AA margin smooth, double dorsal keel. Medial - - - - - Z part of body broadest. With one long toe, and - - - - Y X with small substilis in both sides. Trophi asym- - - - - X X metric; left manubria hook-shaped. Left uncus - - - - - longer. Known from Malaysia, Singapore, and W South America. - - - - - V ) for Central America region. America Central for )

• Trichocerca hollaerti De Smet, 1990 (Fig. - - - - U X 11): Dorsal keel covers all body. Head aperture - - - - T X margins with longitudinal folds. Right toe elon------S gated, the left one “S”-like. It has been reported - - - - -

R in Africa, and Brazil (Segers and Sarma 1993). - - - - - Q - - - - - P - - - - - DISCUSSION O - - - - - Ñ In total 47 taxa are new to the entire - - - - - N Central America region (Collado et al. 1984, De - - - - - M Ridder and Segers 1992). Most of these species - - - - L X are known in southern Mexico (Sarma 1999, - - - - -

K García-Morales and Elías-Gutiérrez 2004). Some species of Euchlanis, Cephalodella, - - - - - J Scaridium and Sinantherina could not be iden- I - - - - - tified due to taxonomic problems or by their - - - - - H poor preservation. Some unidentified bdelloids - - - - - G TABLE 2 (Continued) TABLE were also present. These doubtful taxa were not List of species and localities - - - - - F included in the list. - - - - - As recorded herein, the fauna from E Guatemala and Belize represents the first con- - - - - D X temporaneous addition since Collado et al. - - - - - C (1984). Thus, the number of new records in - - - - B X the list is not surprising. Guatemala and Belize - A X X X X are small countries in size, but they possess

• complex and varied environments as highlands

• and lowlands with rainforest, floodplains, man-

• groves, savanna, Pinus forest, etc. (Archibold 1995). In particular, the region of Pine Ridge, located in the Mayan mountains, has a dramatic change from tropical vegetation to a Pinus for- (Hauer, 1953)* (Hauer, + est in just few meters, possibly as a result of (Rousselet, 1893)+ (Hudson, 1875)+ Shiel & Sanoamuang, 1993* + 1993* Sanoamuang, & Shiel changes in the soil composition and altitude (Levander, 1892)* (Levander, (Elías-Gutiérrez et al. 2006). Here we found the lower temperatures of water and several unique records, as L. rhomboidula. For comparison, small countries such as Ecuador and Bolivia have reports of ca. 300 Conochilus dossuarius Hexarthra fennica H. intermedia f. braziliensis Filinia novaezelandiae novaezelandiae Filinia Collotheca cf. pelagica Species 17. Family Conochilidae 18. Family Hexarthridae 19. Family Filiniidae 20. Family Collothecidae A-K belong to Guatemala and L-AB belong to Belize. For complete names of the localities see Table 1. New records: (*) for Belize; (+) for Guatemala; ( Guatemala; for (+) Belize; for (*) records: New 1. Table see localities the of names complete For Belize. to belong L-AB and Guatemala to belong A-K and 153 species recorded in their freshwater

578 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 55 (2): 569-584, June 2007 Fig. 2. E. semicarinata (dorsal). Fig. 3A. L. apsicora (dorsal), B. Ventral. Fig. 4A. L. cryphaea (dorsal), B. Ventral. systems, respectively (Koste and Böttger 1992, of some rotifers recorded earlier from Africa, Segers et al. 1998). Asia, Europe, New Zealand, USA, and South Out of the 119 taxa reported here, up to America (De Ridder and Segers 1992). For 70 % are cosmopolitan, about 6 % are tropico- example this is the first time that L. rhomboid- politans, 4.2 % are subtropical and 6.7 % have ula is reported in the Americas. limited distribution to one locality. Two taxa Systems with highest and lowest diversity (1.7 %) are neotropical: Brachionus havana- in Guatemala were Petén-Itza and Raxruja with ensis and Hexarthra intermedia f. braziliensis. 53 and 3 species, respectively. In Belize, the The present study demonstrated the occurrence localities La Democracia and Blue Hole had

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 55 (2): 569-584, June 2007 579 Fig. 5. L. rhomboidula (ventral). Fig. 6A. L. curvicornis f. lofuana (ventral), B. Close view of foot. Fig. 7A. L. whitfordi (dorsal), B. Ventral.

Fig. 8. A. M. maculata (habitus ventral), B. trophi (ventral). Fig. 9. S. bostjani (lateral). Fig. 10. T. elongata f. braziliensis (lateral). Fig. 11. T. hollaerti (ventral).

580 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 55 (2): 569-584, June 2007 49 and 6 taxa, respectively. These numbers tricocercids. The cluster analyses confirmed are low, when compared with other neotropi- this statement. We found a major group (letter cal systems. For example Segers et al. (1998) “A” in dendrogram, Fig. 12) formed by the found 104 species of rotifers in only three Southeast Mexico, Yucatan Peninsula, Belize samples from Laguna Bufeos, Bolivia and and Guatemala. As we stated, here we found Sarma and Elías-Gutiérrez (1998) found 78 the majority of rotifer species with South species in a Mexican pond. The majority of American affinity, emphasizing the transitional the species recorded in both, Petén-Itza and La nature of this region between the Nearctics Democracia, were present in littoral samples and Neotropics. North of Mexico has more collected from submerged vegetation, which affinity with the Nearctic fauna, with species harbors a high diversity (Uicab-Sabido 2001). as Keratella paludosa (Lucks), Itura chamadis It should be noticed that all samples from Harring and Myers, Ascomorphella volvocicola Petén-Itza Lake were taken in the west side, (Plate) and Synchaeta tremula (O.F. Müller) near the Flores Island and San Benito town, (García-Morales 2001), and the Pacific coastal the most affected region by human pollution. slope denote influence of species more toler- The east shore, less affected, was not sampled ant to salinity. The “A” group also shows clear in this survey. differences between both Mexican southern Knowledge on the rotifer fauna of Central regions, for example, in Yucatan Peninsula America is still limited when compared with there are more taxa related to alkaline waters that of North and South America. Presently, (García-Morales and Elías-Gutiérrez 2004), Panama is the country with the highest number than in the rest of the Southeast. Belize and of species recorded (180 taxa) in the region (De Ridder and Segers 1992). Since the early report by Harring 1914, who recorded 109 species, 71 taxa have been added (Koste and De Paggi 1982, Collado et al. 1984). With this study the number of species of Guatemala and Belize increased, but remains still behind that of Panama. The necessity of system- atic research on rotifers is urgent in the rest of Central American countries. For example, in Nicaragua there are recorded 52 species only, Costa Rica 41, Honduras 24, and El Salvador 13 (De Ridder and Segers 1992). Central America is an important biogeo- graphic region; it constitutes a bridge (since Pliocene/Pleistocene age) between North and South America (Cox and Moore 1993). The region shows a mixture of species (De Ridder and Segers 1992); nevertheless, the overall influence of the neartic fauna is weak (Proalides tentaculatus, Keratella tauroceph- ala, Lepadella triba and L. whitfordi) (De Ridder and Segers 1992). A good number of taxa have clear affinities with that of South Fig. 12. Cluster analyses of the rotifers in Belize, Guatemala America and the lowlands of the east coast and adjacent Mexico. 1. Yucatan Peninsula; 2. Southeast limit their distributional patterns. This fauna Mexico; 3. Belize; 4. Guatemala; 5. Mexican Central is formed mainly by lecanids, lepadelids and Plateau; 6. North 7. Pacific slope.

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 55 (2): 569-584, June 2007 581 Guatemala form other sub-group, but with AS019) for the study of the zooplankton in some differences: as seen in Table 1, a high southeast Mexico. number of taxa represented in Belize (37 spe- cies) are not present in Guatemala, and vice versa 16 species are only found in the latter. RESUMEN Nevertheless, the whole community makes both countries to form a cluster, separated El análisis de muestras procedentes del norte de from the south of Mexico. Regardless of the Guatemala y centro-sur de Belice, recolectadas en marzo de 2000 y junio de 2001, dio como resultado la presencia few collecting localities in both countries (11 de 119 especies. Se presenta una breve descripción de in Guatemala and 18 in Belize), the rotifers diez taxones seleccionados con base en sus distribuciones appear to show a peculiar assemblage related restringidas en ciertos ámbitos de América y el viejo to particular environments across this tran- continente: Euchlanis semicarinata, Lepadella apsicora, sitional zone that should be confirmed with L. cryphaea, Lecane curvicornis f. lofuana, L. whitfordi, Monommata maculata, Scaridium bostjani, Trichocerca further studies. elongata f. braziliensis, y T. hollaerti. Por primera vez Our results and the analysis of other records se informa Lepadella rhomboidula en el continente suggest that the tropical forest stretching from americano. Las especies registradas incluyen 71 % con the Amazon River basin to Central America distribución cosmopolita y 6 % tropicopolitas; 4.2 % se and Southeastern Mexico is the route by which encuentran restringidas a los subtrópicos. La máxima y mínima diversidad de especies en Guatemala corres- the South American fauna penetrated north- pondió al lago Petén-Itza (con 53 taxones) y al estanque wards (Cox and Moore 1993). This seems to Raxruja (con tres). El estanque La Democracia (con 49 be true for rotifer fauna, as well as other groups taxones) y el Blue Hole (con seis) fueron máximo y míni- of aquatic invertebrates as cladocerans (Elías- mo en Belice. Del total encontrado, 47 especies se regis- Gutiérrez et al. 2001, 2006), and few calanoid tran por primera vez en Centroamérica. Una comparación entre la fauna de rotíferos de estos dos países y México copepods as Prionodiaptomus colombiensis refleja una similitud; sin embargo, también se aprecian Thiébaud (Suárez-Morales et al. 2005). ciertas diferencias que permiten establecer un gradiente Indeed, particular environments from que va de la región neotropical hacia la neártica. Guatemala (as intermittent rivers, and sink- holes in Alta Verapaz) and Belize (the Mayan Palabras clave: rotífero, taxonomía, diversidad, distribu- mountains, and many cave-related systems) ción, Guatemala, Belice. deserve more intensive sampling in different seasons of the year to uncover more details about their peculiar microfauna. REFERENCES

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Segers, H., N.L. Ferrufino & L. De Meester. 1998. Diversity REFERENCIAS DE INTERNET and zoogeography of Rotifera (Monogononta) in a flood plain lake of the Ichilo river, Bolivia, with Jersabek, C.D., H. Segers & P.J. Morri. 2003. An illustrated notes on little-know species. Int. Rev. Hydrobiol. 83: online catalogue of the Rotifera in the Academy 439-448. of Natural Sciences of Philadelphia (version 1.0: 2003-April-8). The Academy of Natural Sciences, Silva-Briano, M. & H. Segers. 1992. Una nueva especie Filadelfia, EEUU. (Available on line: http://data. del género Brachionus (Rotifera, Monogononta) del acnatsci.org/biodiversity_databases/rotifer.php).

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